The question of whether sauropod cervicals got longer through ontogeny came up in the comment thread on Mike’s “How horrifying was the neck of Barosaurus?” post, and rather than bury this as a comment, I’m promoting it to a post of its own.

The short answer is, yeah, in most sauropods, and maybe all, the cervical vertebrae did lengthen over ontogeny. This is obvious from looking at the vertebrae of very young (dog-sized) sauropods and comparing them to those of adults. If you want it quantified for two well-known taxa, fortunately that work was published 16 years ago – I ran the numbers for Apatosaurus and Camarasaurus to see if it was plausible for Sauroposeidon to be synonymous with Pleurocoelus, which was a real concern back in the late ’90s (the answer is a resounding ‘no’). From Wedel et al. (2000b: pp. 368-369):

Despite the inadequacies of the type material of Pleurocoelus, and the uncertainties involved with referred material, the genus can be distinguished from Brachiosaurus and Sauroposeidon, even considering ontogenetic variation. The cervical vertebrae of Pleurocoelus are uniformly short, with a maximum EI of only 2.4 in all of the Arundel material (Table 4). For a juvenile cervical of these proportions to develop into an elongate cervical comparable to those of Sauroposeidon, the length of the centrum would have to increase by more than 100% relative to its diameter. Comparisons to taxa whose ontogenetic development can be estimated suggest much more modest increases in length.

Carpenter & McIntosh (1994) described cervical vertebrae from juvenile individuals of Apatosaurus and Camarasaurus. Measurements and proportions of cervical vertebrae from adults and juveniles of each genus are given in Table 4. The vertebrae from juvenile specimens of Apatosaurus have an average EI 2.0. Vertebrae from adult specimens of Apatosaurus excelsus and A. louisae show an average EI of 2.7, with an upper limit of 3.3. If the juvenile vertebrae are typical for Apatosaurus, they suggest that Apatosaurus vertebrae lengthened by 35 to 65% relative to centrum diameter in the course of development.

The vertebrae from juvenile specimens of Camarasaurus have an average EI of 1.8 and a maximum of 2.3. The relatively long-necked Camarasaurus lewisi is represented by a single skeleton, whereas the shorter-necked C. grandis, C. lentus, and C. supremus are each represented by several specimens (McIntosh, Miller, et al. 1996), and it is likely that the juvenile individuals of Camarasaurus belong to one of the latter species. In AMNH 5761, referred to C. supremus, the average EI of the cervical vertebrae is 2.4, with a maximum of 3.5. These ratios represent an increase in length relative to diameter of 30 to 50% over the juvenile Camarasaurus.

If the ontogenetic changes in EI observed in Apatosaurus and Camarasaurus are typical for sauropods, then it is very unlikely that Pleurocoelus could have achieved the distinctive vertebral proportions of either Brachiosaurus or Sauroposeidon.

C6 of Apatosaurus CM 555 – despite having an unfused neural arch and cervical ribs, the centrum proportions are about the same as in an adult.

A few things about this:

From what I’ve seen, the elongation of the individual vertebrae over ontogeny seems to be complete by the time sauropods are 1/2 to 2/3 of adult size. I get this from looking at mid-sized subadults like CM 555 and the hordes of similar individuals at BYU, the Museum of Western Colorado, and other places. So to get to the question posed in the comment thread on Mike’s giant Baro post – from what I’ve seen (anecdata), a giant, Supersaurus-class Barosaurus would not necessarily have a proportionally longer neck than AMNH 6341. It might have a proportionally longer neck, I just haven’t seen anything yet that strongly suggests that. More work needed.

Juvenile sauropod cervicals are not only shorter than those of adults, they also have less complex pneumatic morphology. That was the point of the figure at the top of the post. But that very simple generalization is about all we know so far – this is an area that could use a LOT more work.

I’ve complained before about papers mostly being remember for one thing, even if they say many things. This is the canonical example – no-one ever seems to remember the vertebrae-elongating-over-ontogeny stuff from Wedel et al. (2000b). Maybe that’s an argument for breaking up long, kitchen-sink papers into two or more separate publications?

This was great: it gave us the opportunity to handle those review comments and get the manuscript turned around into an already-reviewed formal journal submission in less then a month from the original talk.

So of course what we did instead was: nothing. For three years.

I can’t excuse that. I can’t even explain it. It’s not as though we’ve spent those three years churning out a torrent of other awesome papers. We’ve both just been … a bit lame.

Anyway, here’s a story that will be hauntingly familiar. A month ago, full of enthusiasm after speaking about Barosaurus at the Liverpool SVPCA, Matt and I found ourselves keen to write up that talk in double-quick time. It’s an exciting tale of new specimens, reinterpretation of an important old specimen, and a neck eight times as long as that 0f a world-record giraffe.

But it would be crazy to write the new Barosaurus paper without first having dealt with the old Barosaurus paper. So now, finally, three years on, we’ve done that. Version 2 of the preprint is now available (Taylor and Wedel 2016), incorporating all the fine suggestions of the people who reviewed the first version — and with a slightly spiffed-up title. What’s more, the new version has also been submitted for formal peer-review. (In retrospect, I can’t think why we didn’t do that when we put the first preprint up.)

A big part of the purpose of this post is to thank Emanuel Tschopp, Mark Robinson, Andy Farke, John Foster and Mickey Mortimer for their reviews back in 2013. I know it’s overdue, but they are at least all acknowledged in the new version of the manuscript.

Now we cross our fingers, and hope that the formally solicited reviews for the new version of the manuscript are as helpful and constructive as the reviews in that first round. Once those reviews are in, we should be able to move quickly and painlessly to a formally published version of this paper. (I know, I know — I shouldn’t offer such a hostage to fortune.)

Meanwhile, I will finally be working on handling the reviews of this other PeerJ submission, which I received back in October last year. Yes, I have been lax; but I am back in the saddle now.

References

Taylor, Michael P., and Mathew J. Wedel. 2016. The neck of Barosaurus: longer, wider and weirder than those of Diplodocus and other diplodocines. PeerJ PrePrints 1:e67v2 doi:10.7287/peerj.preprints.67v2

May 17, 2016

UPDATE 19 May 2016

I belatedly realized that I caused some confusion in the original version of this post. This will hopefully sort things out:

The ventrolateral processes (1) are nothing new. As Ken Carpenter pointed out in a comment, Hatcher noted them back in 1901 in his monograph on Diplodocus carnegii. These are the features I describe below as being, “huge in Barosaurus, big in Diplodocus, small in Apatosaurus, and nonexistent in Haplocanthosaurus, Camarasaurus, and the brachiosaurids, at least from what I’ve seen.” To clarify: occasionally in camarasaurs and frequently in brachiosaurs you can trace a ridge along the ventrolateral margin of the centrum from the parapophysis to the cotyle. But these ridges are basically just the ‘corners’ of the centrum, leftover by the lateral and ventral waisting of the centrum – they do not project beyond the margin of the cotyle. In contrast, what I’ve been calling the ventrolateral flanges in diplodocids do project beyond the margins of the cotyle – they are additive structures, not just architectural leftovers. They also don’t vary much, other than to be more pronounced in more posterior cervicals.

The irregular ventral ridges (2) are a totally different thing. They’re on or near the sagittal midline of the centrum, usually restricted to the anteroposterior middle of the ventral centrum (so, about halfway between the condyle and the cotyle), and as my preferred term implies, highly variable among individuals and even among vertebrae in a series.

Hope that helps! (Original post starts below.)

– – – – – – – – – – – – – – – – – – – – –

Back in 2005 I visited BYU while I was working on my dissertation. Back then I noted ventral ridges in a few diplodocine cervical vertebrae. (I hesitate to call such flimsy things ‘keels’.)

Up above is BYU 16918, a mid-to-posterior cervical vertebra of Diplodocus from the famous Dry Mesa Quarry. Here it is again in posterior view:

The things I have labeled VLF here are ventrolateral flanges, which are huge in Barosaurus, big in Diplodocus, small in Apatosaurus, and nonexistent in Haplocanthosaurus, Camarasaurus, and the brachiosaurids, at least from what I’ve seen. See this post for details. I know that the left VLF here looks like a second ridge, but the cotyle is broken off in such a way that we’re seeing the fossa just dorsal to the VLF margin. The ridge itself is skewed to the right, which could be natural or a result of taphonomy – as you can see from the photo at the top of the post, this vert has seen better days.

Here’s another Dry Mesa vert, BYU 11617, this time an anterior cervical of Barosaurus and in left lateral view:

Again in right lateral view – on this side you can see the fossa in the VLF more clearly:

And here’s the ventral view showing the ridge:

I noted these things in my notebook back when, filed them under, “Huh. How about that?” and went on with life.

Then last week Mike and I were at the North American Museum of Ancient Life in Lehi, Utah, and we saw this super-nice Barosaurus cervical on display in the prep lab (left ventro-lateral view). Check out the monster ventrolateral flanges, and the ridges between them at about mid-centrum.

Here’s another view, a more square-on ventral this time:

We owe a big thank you to Rick Hunter, who let us into the prep lab at the North American Museum of Ancient Life to see the Barosaurus material up close.

So what’s the deal with these ridges? I assume that they’re caused by pneumatic diverticula remodeling the ventral surface of the centrum. We know that such diverticula were down there because there are actual foramina on the ventral centrum in Supersaurus, many apatosaurines (Lovelace et al., 2008), many brachiosaurids, and probably loads of other things that haven’t been checked. Oddly enough, I’ve never seen the ridges in any of those other taxa. It seems that you get foramina or ridges, but not both. I have no idea what’s up with that – to paraphrase Neal Stephenson, Barosaurus cervicals are confections of air and marketing, and you’d think that if any sauropod would have straight-up foramina down there, it would be Barosaurus. But Barosaurus gets ridges and clunky old Apatosaurus gets foramina (sometimes, not all the time).

May 12, 2016

Things remain frantic on the Sauropocalypse tour. Today, we were back at the BYU Museum of Paleontology, working on four or five separate projects. Here’s Matt, photographing broken bone of the iconic Supersaurus cervical BYU 9024, while a pallet of Big Pink Apatosaur cervicals wait for attention in the background:

You’ve seen this bone before – I first posted on it 8 years ago this month, and it turned up again here and here. It is still the longest known vertebra of any animal that has ever lived.

And here’s Mike, getting Jensen’s sculpture of the same vertebra down from storage to compare it to the original:

In Jensen’s (1985) original description of this vertebra – which he at first referred to Ultrasauros – the only relevant illustration he included was one of the model, so it was good to see this bit of history in the flesh (Jensen did include photos of the actual bone in later papers). We’ll show the two vertebrae, real and sculpted, side by side in a future post.

References

Jensen, J. A. 1985. Three new sauropod dinosaurs from the Upper Jurassic of Colorado. Great Basin Naturalist 45, 697-709.

May 9, 2016

Not much to say this time – the pictures tell the story for now.

It was a pretty transcendental experience, as I imagine it must be for anyone who loves dinosaurs, or has a pulse.

A huge thank-you to Dan Chure, the Park Paleontologist for the Monument, who conveyed us safely up and down the Wall, taught us about the prehistory of the site and the human history of its excavation and conservation, held scale bars, moved backpacks, took photos, and generally seemed to be having just as much fun as we were. This has been a common theme on the trip – every single person we’ve interacted with at a museum or fossil site has been unfailingly welcoming and generous with their time and knowledge. Whatever challenges vert paleo faces, a lack of wonderful people is not one of them.

I was up there, too, for the second time in my life – that will be a post for another day. For now, just bask in the glory of Mike basking in the glory of a literally mind-numbing array of amazing fossils.

May 7, 2016

At the Prehistoric Museum in Price, Utah, our host Ken Carpenter invited us to jump right into the Camarasaurus pit and start pulling apart their beautiful specimen. We did. Here is Matt, looking as happy as I’ve ever seen him:

The pit is the central exhibit of the museum’s palaeontology hall. You can look down on its Jurassic scene from the balcony above:

Theres a very nice Stegosaurus and an Allosaurus in pursuit of some kind of ornithopod, but needless to say the star of the show is the dead Camarasaurus that lies on the ground, well associated but partially articulated.

It’s a beautifully undistorted specimen, and we were amazed and delighted when Ken not only gave us permission to hop over the barrier and get closer to it, but even to move the elements around to better measure and photograph them. We spend the morning with the baby, concentrating on four anterior cervicals, and could happily have spent much, much longer.

A shot across the room at ground level:

Further bulletins as and when we find time to post. Can’t write more now, we’re off to the big wall of awesome at Dinosaur National Monument!

May 3, 2016

When I was back in Oklahoma in March, I met with Anne Weil to see some of the new Apatosaurus material she’s getting out of her Homestead Quarry. It’s nice material, but that’s a post for another day. Anne said something that really resonated with me, which was, “I love it when you guys post about vertebral morphology, because it helps me learn this stuff.” Okay, Anne, message received. This will begin to make things right.

I spent a week at BYU back in 2005, collecting data for my dissertation. One of the first things I had to do was teach myself how to identify the vertebrae of different sauropods, because BYU has just about all of the common Morrison taxa. These are the notes I made back then.

I always planned to do something with them – clean them up, get them into a more usable form. There are a lot of scribbly asides that are probably hard for others to read, and it would be more useful if I put the easily confused taxa next to each other – Barosaurus next to Brachiosaurus, for example. And I didn’t go into serial changes at all.

Still, hopefully someone will find these useful. If there are things I missed or got wrong, the comment thread is open. And if you want all four spreads in one convenient package, here’s a PDF: Wedel 2005 notes on Morrison sauropod cervicals